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Biuletyn Sekcji Dipterologicznej Polskiego Towarzystwa Entomologicznego DIPTERON

Bulletin of the Dipterological Section of the Polish Entomological Society

ISSN 1895–4464 Tom 32: 77-93

Akceptacja: 22.12.2016 Wrocław 31 XII 2016

Biting midges (Diptera: Ceratopogonidae) of Poland: a checklist

Wykaz kuczmanów Polski (Diptera: Ceratopogonidae)

RYSZARD SZADZIEWSKI & PATRYCJA DOMINIAK*

Department of Invertebrate Zoology and Parasitology, University of Gdańsk

Wita Stwosza 59, 80-308 Gdańsk, Poland

*Corresponding author email: heliocopris@gmail.com

ABSTRACT. With 220 species of biting midges in 21 genera, this checklist includes 13 species

more than the previous one. Haematophagous biting midges feeding on mammalian and avian

blood include 52 species from the genus Culicoides and one from Forcipomyia. Ceratopogon

naccinervis BORKENT, in BORKENT & WIRTH 1997 is recorded for the first time in the Polish

fauna.

KEY WORDS: faunistics, Poland, nomina dubia, new record

INTRODUCTION

Biting midges (Diptera: Ceratopogonidae) are small nematocerous dipterans that are

common in all kinds of aquatic, semiaquatic and moist terrestrial habitats worldwide. The

apneustic larvae of biting midges are easily distinguished from chironomids by the heavily

sclerotized, unique pharyngeal apparatus located in the middle of the head capsule. Adult

biting midges have two medial wing veins M1 and M2 and relatively short fore legs; females

are usually armed with functional biting mouthparts. The antennae of males usually have long

appressed plumose setae. In contrast, the median vein in adult Chironomidae is always single,

the fore legs are usually long, the female mouthparts greatly reduced and the male antennae

have perpendicular plumose setae (SZADZIEWSKI 2007).

Larvae of biting midges are common inhabitants of many types of aquatic and semiaquatic

habitats. They live mostly in shallow waters of lakes, ponds, rivers, streams, springs, pools as

well as wet margins of inland water bodies, marshes, pit bogs, wet meadows, mosses and

matts of algae on soil, rocks or floating in water. In rivers largely occur the same groups of

biting midges which live in lakes and ponds. It is believed that only in the subgenus

Atrichopogon (Psammopogon) some rheophilic species occurring exclusively in cold streams

exsist (NIELSEN 1951). Biting midges of the genera Culicoides, Dasyhelea, Forcipomyia,

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Atrichopogon and Bezzia are also common in saline inland or coastal habitats. For example,

Culicoides longicollis is an aquatic halobiont, whose larvae live in filamentous algae of

strongly mineralized inland small water bodies in Central Poland (SZADZIEWSKI 1983). Many

biting midge species of the genera Forcipomyia and Atrichopogon and some Culicoides live

in typical terrestrial habitats such as rotting wood, plant debris, under bark, in macrofungi, in

cow and horse dung, rotting fungi and ant nests. The larvae of Dasyhelea flavifrons,

Culicoides semimaculatus and C. flavipulicaris are typical inhabitants of tree holes and sap

exuding from various trees (GLUKHOVA 1989, DOMINIAK 2012), although the latter species is

also known to occur in mud rich in organic matters and watering channels (USLU & DIK

2007).

The larvae of the subfamily Ceratopogoninae are mostly predaceous, consuming eggs and

larvae of chironomids, mosquitoes, trichopterans, nematodes, rotifers, annelids, protozoans

and other small invertebrates. They also feed on microalgae, dead insect larvae, fungi, detritus

and bacteria. The larvae of Dasyheleinae and Forcipomyiinae are saprophagous: they eat

decomposed organic plant and fungal debris, as well as filamentous algae and diatoms

(SZADZIEWSKI 2007).

The feeding habits of adult biting midges are highly diversified. Haematophagy and

completely functional mouthparts are known only in females. Males feed only on water and

sugar in flower nectar and honeydew. Both sexes of all species of Dasyhelea and many of

Forcipomyia feed exclusively on flower nectar. Female biting midges consume nectar as a

supplementary protein meal necessary for egg development. Some haematophagous species

are capable of ovipositing the first clutch of eggs without feeding on blood (autogeny) but do

require a protein meal to initiate subsequent ovarian cycles (GLUKHOVA 1989).

Females of Culicoides and Forcipomyia (Lasiohelea) sibirica (BUYANOVA, 1962) feed on

the blood of mammals and birds and are often serious pests, especially in mountain areas

(GLUKHOVA 1989). Another species of Lasiohelea, i.e. F. (L.) velox (WINNERTZ, 1852), is a

bloodsucking parasite attacking amphibians and spreading a filarial nematode Icosiella

neglecta (DIESING, 1851) among frogs (SZADZIEWSKI 1986). Haematophagous biting midges

are vectors of pathogenic arboviruses, filarial parasites and protozoan blood parasites in frogs,

birds and mammals. Some species of Forcipomyia and Atrichopogon are parasites of the

larvae and adults of larger insects (butterflies, beetles, neuropterans), feeding on their

haemolymph. Biting midges of the subgenus Atrichopogon (Meloehelea) are

cantharidophilous as they feed on the haemolymph of beetles producing cantharidin. Females

of Atrichopogon pavidus are pollinivorous as they obtain proteins and fat-rich food from the

pollen grains of honeysuckle (Lonicera). Females of other species of the subfamily

Ceratopogoninae, apart from Culicoides, are predators on the adults of small insects, usually

male chironomids and small Ephemeroptera. They inject into their prey a saliva rich in

proteolytic enzymes which dissolves tissues, and then suck out the resulting digested fluid.

Within the tribe Sphaeromiini, females often kill and eat males during copulation

(SZADZIEWSKI et AL. 1997, BORKENT et AL. 2009).

Ceratopogonidae belong to the group of insects with a well-known evolution based on a

rich fossil record (SZADZIEWSKI 1996), while the recent World fauna is also quite reach and

comprises over 6 000 extant species (BORKENT 2016). Almost 600 valid species names have

been reported from Europe (SZADZIEWSKI et AL. 2013).

The previous checklist of the Polish fauna contained 207 species (SZADZIEWSKI 2007). To

that list we have added species from the following genera: Alluaudomyia (SZADZIEWSKI et AL.

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2015, one species described from Poland as new to science), Atrichopogon (DOMINIAK et AL.

2007), Bezzia (DOMINIAK & SZADZIEWSKI 2008), Ceratopogon (DOMINIAK & SZADZIEWSKI

2008, present record), Culicoides (DOMINIAK & SZADZIEWSKI 2008, KIRKEBY & DOMINIAK

2014), Dasyhelea (DOMINIAK & SZADZIEWSKI 2008), Forcipomyia (DOMINIAK &

MICHALCZUK 2009, ALWIN & SZADZIEWSKI 2013, DOMINIAK et AL. 2015), Monohelea

(DOMINIAK & MICHALCZUK 2009) and Stilobezzia (ALWIN & SZADZIEWSKI 2012).

The present checklist of the biting midges of Poland includes 220 species in 21 genera. The

species list still contains two doubtful names: Atrichopogon longiserrus (KIEFFER, 1921), and

Atrichopogon singularis (KIEFFER, 1921) (nomina dubia), described from Polish Silesia.

Their descriptions are incomplete, and the type specimens were lost during a fire in Budapest

in 1956. Ceratopogon naccinervis BORKENT, in BORKENT & WIRTH 1997 is recorded for the

first time in the Polish fauna. Poland with 220 species of biting midges reported is among the

best studied countries in Europe. More species are reported only from Germany (256) and

France (241). In other relatively well studied countries lower number of species is reported,

for example in Estonia – 184, Czechia – 175, Great Britain – 169, and Slovakia – 137

(SZADZIEWSKI et AL. 2013).

CHECKLIST

Order Diptera LINNAEUS, 1758

Family Ceratopogonidae NEWMAN, 1834

Subfamily Forcipomyiinae LENZ, 1934

Atrichopogon KIEFFER, 1906 (23)

Atrichopogon (Atrichopogon KIEFFER, 1906) (11)

- aethiops (GOETGHEBUER, 1920) (Kempia)

- alveolatus NIELSEN, 1951 (Atrichopogon)

- bargaensis REMM, 1972 (Atrichopogon)

- brunnipes (MEIGEN, 1804) (Ceratopogon)

Syn.: longinervis KIEFFER, 1919; semiensis REMM, 1972

- fuscus (MEIGEN, 1804) (Ceratopogon)

Syn.: fuscipes ZETTERSTEDT, 1850; fossicola KIEFFER, 1922

- infuscus GOETGHEBUER, 1929 (Atrichopogon)

- longicalcar REMM, 1961 (Atrichopogon)

- longiserrus (KIEFFER, 1921) (Kempia) [nomen dubium]

- minutus (MEIGEN, 1830) (Ceratopogon)

Syn.: fortiserrus KIEFFER, 1924; turficola KIEFFER, 1925; parviforceps TOKUNAGA, 1940

- pavidus (WINNERTZ, 1852) (Ceratopogon)

Syn.: pollinivorus DOWNES, 1955

- singularis (KIEFFER, 1921) (Kempia) [nomen dubium]

Atrichopogon (Lophomyidium CORDERO, 1929) (2)

- fusculus (COQUILLETT, 1901) (Ceratopogon)

Syn.: polydactylus NIELSEN, 1951

- rostratus (WINNERTZ, 1852) (Ceratopogon)

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Syn.: transversalis KIEFFER, 1918; ventralis KIEFFER, 1918; homopterus KIEFFER, 1919; coracellus

KIEFFER, 1919; nigriventris KIEFFER, 1919; putredinis KIEFFER, 1922

Atrichopogon (Meloehelea WIRTH, 1956) (5)

- lucorum (MEIGEN, 1818) (Ceratopogon)

Syn.: sylvaticus WINNERTZ, 1852; setosipennis KIEFFER, 1911; bidentatus KIEFFER, 1924; seminitidus

GOETGHEBUER, 1947

- meloesugans KIEFFER, 1922 (Atrichopogon)

- oedemerarum STORÅ, 1939 (Atrichopogon)

- orbicularis KIEFFER, 1919 (Atrichopogon)

- winnertzi GOETGHEBUER, 1922 (Atrichopogon)

Syn.: torgnyensis GOETGHEBUER, 1949

Atrichopogon (Psammopogon REMM, 1979) (2)

- flaveolus ZILAHI-SEBESS, 1936 (Atrichopogon)

- muelleri (MÜLLER, 1905) (Ceratopogon)

Syn.: muelleri KIEFFER, 1906; cornutus NIELSEN, 1951; globularis MAYER, 1934

Atrichopogon (Psilokempia ENDERLEIN, 1936) (3)

- appendiculatus (GOETGHEBUER, 1920) (Kempia)

Syn.: armativentris KIEFFER, 1923; adjacens KIEFFER, 1924

- forcipatus (WINNERTZ, 1852) (Ceratopogon)

Syn.: silesiacus KIEFFER, 1919; hamifera GOETGHEBUER, 1920

- maculatus (LUNDSTRÖM, 1910) (Ceratopogon)

Syn.: avastensis REMM, 1959; hamulatus REMM, 1971

Forcipomyia MEIGEN, 1818 (38)

Forcipomyia (Caloforcipomyia SAUNDERS, 1956) (1)

- glauca MACFIE, 1934 (Forcipomyia)

Syn.: monilis GOETGHEBUER, 1934; diversipes GOETGHEBUER, 1936; splendida WIRTH, 1951

Forcipomyia (Euprojoannisia BRÉTHES, 1914) (5)

- alacris (WINNERTZ, 1852) (Ceratopogon)

- borealis REMM, 1966 (Forcipomyia)

- palustris (MEIGEN, 1804) (Ceratopogon)

Syn.: hirtipennis MALLOCH, 1915; turfacea KIEFFER, 1925

- phlebotomoides BANGERTER, 1933 (Forcipomyia)

Syn.: subsaltans REMM, 1961

- titillans (WINNERTZ, 1852) (Ceratopogon)

Syn.: divaricatus WINNERTZ, 1852; maricarmenae SAHUQUILLO & GIL COLLADO, 1982

Forcipomyia (Forcipomyia MEIGEN, 1818) (17)

- altaica REMM, 1972 (Forcipomyia)

- bipunctata (LINNAEUS, 1767) (Tipula)

Syn.: trichopterus MEIGEN, 1804; ambigua MEIGEN, 1804; geniculatus GUÉRIN, 1833; palmensis

SANTOS ABREU, 1918; squamaticrus KIEFFER, 1919; tenuisquama KIEFFER, 1924; laguncula KIEFFER,

1925; flavipubens GOETGHEBUER, 1927

- brevipennis (MACQUART, 1826) (Ceratopogon)

Syn.: lateralis BOUCHÉ, 1834; specularis COQUILLETT, 1901; coprophila KIEFFER, 1914; disticta

KIEFFER, 1919; hirtidorsum KIEFFER, 1919; nigrimanus KIEFFER, 1919; bifilis KIEFFER, 1922; subnigra

TOKUNAGA, 1940

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- ciliata (WINNERTZ, 1852) (Ceratopogon)

Syn.: brunnipes PERRIS, 1847; perrisi KIEFFER, 1901; boleti KIEFFER, 1901; turfosa KIEFFER, 1925;

canicularis GOETGHEBUER, 1948; zlatensis DAMIAN-GEORGESCU, 1972

- costata (ZETTERSTEDT, 1838) (Ceratopogon)

Syn.: piceus WINNERTZ, 1852; latipalpis KIEFFER, 1901; meinerti KIEFFER, 1915; corticicola KIEFFER,

1919; turficola KIEFFER, 1925

- hygrophila KIEFFER, 1925 (Forcipomyia)

Syn.: sphagnicola KIEFFER, 1925; occidentalis WIRTH, 1952

- kaltenbachi (WINNERTZ, 1852) (Ceratopogon)

- nigra (WINNERTZ, 1852) (Ceratopogon)

Syn.: sphagnorum KIEFFER, 1925

- nigrans REMM, 1962 (Forcipomyia)

- pallida (WINNERTZ, 1852) (Ceratopogon)

- pulchrithorax EDWARDS, in SAUNDERS 1924 (Forcipomyia)

- radicicola EDWARDS, in SAUNDERS 1924 (Forcipomyia)

Syn.: padi REMM, 1979

- regula (WINNERTZ, 1852) (Ceratopogon)

Syn.: metatarsis TOKUNAGA, 1940; longimaculata TOKUNAGA, 1940

- sahariensis KIEFFER, 1923 (Forcipomyia)

Syn.: armaticrus KIEFFER, 1923; tuzeti HUTTEL & HUTTEL, 1952; acanthophora REMM, in HAVELKA

1976; onusta REMM, 1980

- sphagnophila KIEFFER, 1925 (Forcipomyia)

Syn.: solonensis WIRTH, 1951; oreophila REMM, 1972

- squamigera KIEFFER, in THIENEMANN & KIEFFER 1916 (Forcipomyia)

Syn.: apricans (KIEFFER, 1919)

- tenuis (WINNERTZ, 1852) (Ceratopogon)

Forcipomyia (Lasiohelea KIEFFER, 1921) (2)

- sibirica (BUYANOVA, 1962) (Lasiohelea)

- velox (WINNERTZ, 1852) (Ceratopogon)

Syn.: halteratus WINNERTZ, 1852; nitens KIEFFER, 1919; pilosipennis KIEFFER, 1919; hungarica

KIEFFER, 1921; decrescens KIEFFER, 1924; silesiae KIEFFER, 1925; montschadskyi DZHAFAROV, 1962

Forcipomyia (Lepidohelea KIEFFER, 1917) (1)

- tibialis REMM, 1961 (Forcipomyia)

Forcipomyia (Microhelea KIEFFER, 1917) (1)

- fuliginosa (MEIGEN, 1818) (Ceratopogon)

Syn.: villosa ZETTERSTEDT, 1850; crudelis KARSCH, 1886; hirtipes DE MEIJERE, 1907; brevimana

LUNDSTRÖM, 1910; inornatipennis AUSTEN, 1912; erucicida KNAB, 1914; crudelis KNAB, 1914;

coquilletti KIEFFER, 1917; tropicus KIEFFER, 1917; obscura SANTOS ABREU, 1918; alboclavatus

KIEFFER, 1919; canaliculata GOETGHEBUER, 1920; nilotheres MACFIE, 1924; ornaticrus INGRAM &

MACFIE, 1924; atripennis GOETGHEBUER, 1935; auripila GOETGHEBUER, 1935; curtimana

GOETGHEBUER, 1935; grisescens GOETGHEBUER, 1935; vicina GOETGHEBUER, 1935; longiradialis

TOKUNAGA, 1940; takagii TOKUNAGA, 1941; wansoni HARANT & BAUR, 1946; brookmani WIRTH,

1952; santosi REMM, 1981

Forcipomyia (Panhelea REMM, 1980) (1)

- aristolochiae (RONDANI, 1860) (Ceratopogon)

Syn.: brevicubitus GOETGHEBUER, 1920; pontica REMM, in REMM & ZHOGOLEV 1968

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Forcipomyia (Pterobosca MACFIE, 1940) (1)

- paludis (MACFIE, 1936) (Pterobosca)

Forcipomyia (Synthyridomyia SAUNDERS, 1956) (3)

- acidicola (TOKUNAGA, 1937) (Lasiohelea)

Syn.: minuta GOETGHEBUER, 1947; colemani WIRTH, 1952

- knockensis GOETGHEBUER, 1938 (Forcipomyia)

Syn.: bequaerti GOETGHEBUER, 1942; abludens REMM, in REMM & ZHOGOLEV 1968

- murina (WINNERTZ, 1852) (Ceratopogon)

Syn.: abdominalis SANTOS ABREU, 1918; aurosparsum KIEFFER, 1919; sulfurea KIEFFER, 1923;

hirtipalpis KIEFFER, 1924; sate KIEFFER, 1925; longitarsis TOKUNAGA, 1940; moascari MACFIE, 1943;

attonsa GOETGHEBUER, 1950; tokunagai WIRTH, 1973; submurina REMM, 1980

Forcipomyia (Thyridomyia SAUNDERS, 1925) (3)

- litoraurea (INGRAM & MACFIE, 1924) (Lasiohelea)

Syn.: minutissima REMM, 1961

- monilicornis (COQUILLETT, 1905) (Ceratopogon)

Syn.: hirta LUNDSTRÖM, 1910; microcera KIEFFER, 1919; palustris SAUNDERS, 1925; curticornis

GOETGHEBUER, 1933; curticornis GOETGHEBUER, 1949; kabashae DE MEILLON, 1959

- rugosa CHAN & LE ROUX, 1970 (Forcipomyia)

Syn.: murinoides REMM, 1971

Forcipomyia (Trichohelea GOETGHEBUER, 1920) (3)

- chaetoptera REMM, 1962 (Forcipomyia)

- eques (JOHANNSEN, 1908) (Ceratopogon)

Syn.: chrysopae MAYER, 1934; chrysopivora TOKUNAGA, 1939

- tonnoiri (GOETGHEBUER, 1920) (Trichohelea)

Syn.: auronitens KIEFFER, 1919; papilionivora EDWARDS, 1923; umbellicola REMM, 1971

Subfamily Dasyheleinae LENZ, 1934

Dasyhelea KIEFFER, 1911 (30)

Dasyhelea (Dasyhelea KIEFFER, 1911) (6)

- bilineata GOETGHEBUER, 1920 (Dasyhelea)

Syn.: insignipalpis KIEFFER, 1925; saxicola EDWARDS, 1929; geleiana ZILAHI-SEBESS, 1931; montana

ZILAHI-SEBESS, 1940; lithotelmatica STRENZKE, 1950; tecticola REMMERT, 1953; dieuzeidei VAILLANT,

1957; karelica GLUKHOVA & BRODSKAYA, 1997

- caesia REMM, 1993 (Dasyhelea)

Syn.: lugensis BRODSKAYA, 1995

- flavifrons (GUÉRIN, 1833) (Ceratopogon)

Syn.: obscurus WINNERTZ, 1852; versicolor WINNERTZ, 1852; dufouri LABOULBÈNE, 1869;

hippocastani MIK, 1888; brevitibialis GOETGHEBUER, 1919; goetghebueri KIEFFER, 1919; lignicola

KIEFFER, 1919; sensualis KIEFFER, 1919; paludicola KIEFFER, 1925; oppressa THOMSEN, 1935;

septuosa BORKENT, 1997

- malleola REMM, 1962 (Dasyhelea)

- pallidiventris (GOETGHEBUER, 1931) (Tetraphora)

Syn.: olivacea REMM, 1962

- stackelbergi REMM, 1993 (Dasyhelea)

Dasyhelea (Dicryptoscena ENDERLEIN, 1936) (6)

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- glukhovae BRODSKAYA, 1996 (Dasyhelea)

- lucida REMM, 1968 (Dasyhelea)

- modesta (WINNERTZ, 1852) (Ceratopogon)

Syn.: aestivus WINNERTZ, 1852; longipalpis KIEFFER, 1913; inclusa KIEFFER, 1918; strobli KIEFFER,

1919; pratensis GOETGHEBUER, 1920; bihamata KIEFFER, 1923; moascari MACFIE, 1943; densipilosa

TOKUNAGA, 1963

- notata GOETGHEBUER, 1920 (Dasyhelea)

Syn.: semistriata GOETGHEBUER, 1921; sziladyi ZILAHI-SEBESS, 1936

- stellata REMM, 1968 (Dasyhelea)

- thienemanni SPĂTARU & DAMIAN-GEORGESCU, 1970 (Dasyhelea)

Dasyhelea (Prokempia KIEFFER, 1913) (3)

- bilobata KIEFFER, 1915 (Dasyhelea)

Syn.: luteiventris GOETGHEBUER, 1934; spiralis REMM, 1966

- dampfi KIEFFER, 1925 (Dasyhelea)

Syn.: turfacea KIEFFER, 1925; estonica KIEFFER, 1925

- flaviventris (GOETGHEBUER, 1910) (Culicoides)

Syn.: halobia (KIEFFER, 1924)

Dasyhelea (Pseudoculicoides MALLOCH, 1915) (14)

- arenivaga MACFIE, 1943 (Dasyhelea)

- auli REMM, 1962 (Dasyhelea)

- bicrenata KIEFFER, 1923 (Dasyhelea)

- bifida ZILAHI-SEBESS, 1936 (Dasyhelea)

Syn.: excellentis BORKENT, 1997

- calycata REMM, 1972 (Dasyhelea)

- corinneae GOSSERIES, 1991 (Dasyhelea)

Syn.: scutellatus MEIGEN, 1830; chonetus YU & ZOU, 2006

- flavoscutellata (ZETTERSTEDT, 1850) (Ceratopogon)

- furva REMM, 1967 (Dasyhelea)

- kyrenica REMM, 1972 (Dasyhelea)

- leptoclada REMM, 1967 (Dasyhelea)

- similaris REMM, 1972 (Dasyhelea)

- turanicola REMM & NAZARMUKHAMEDOV, 1969 (Dasyhelea)

Syn.: serrata NAVAI, 1994

- turficola KIEFFER, 1925 (Dasyhelea)

Syn.: grenieri CLASTRIER, 1966; malibui YU, 2008

- unguistyla REMM, 1972 (Dasyhelea)

Dasyhelea (Sebessia REMM, 1979) (1)

- holosericea (MEIGEN, 1804) (Ceratopogon)

Subfamily Ceratopogoninae NEWMAN, 1834

Tribe Culicoidini KIEFFER, 1911

Culicoides LATREILLE, 1809 (52)

Culicoides (Avaritia FOX, 1955) (6)

- abchazicus DZHAFAROV, 1964 (Culicoides)

- chiopterus (MEIGEN, 1830) (Ceratopogon)

Syn.: amoenus WINNERTZ, 1852; similis GOETGHEBUER, 1927; dobyi CALLOT & KREMER, 1969

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- dewulfi GOETGHEBUER, 1936 (Culicoides)

Syn.: pseudochiopterus DOWNES & KETTLE, 1952

- gornostaevae MIRZAEVA, 1984 (Culicoides)

- obsoletus (MEIGEN, 1818) (Ceratopogon)

Syn.: varius WINNERTZ, 1852; yezoensis MATSUMURA, 1911; obscuripes SANTOS ABREU, 1918;

lacteinervis KIEFFER, 1919; rivicola KIEFFER, 1921; clavatus KIEFFER, 1921; heterocerus KIEFFER,

1921; pegobius KIEFFER, 1922; kabyliensis KIEFFER, 1922; concitus KIEFFER, 1922; intermedius

OKADA, 1941; sintrensis CAMBOURNAC, 1956; seimi SHEVCHENKO, 1967

- scoticus DOWNES & KETTLE, 1952 (Culicoides)

Culicoides (Beltranmyia VARGAS, 1953) (4)

- circumscriptus KIEFFER, 1918 (Culicoides)

Syn.: nadayanus KIEFFER, 1918; edwardsi GOETGHEBUER, 1921; algarum KIEFFER, 1924; salicola

KIEFFER, 1924; pictidorsum KIEFFER, 1924; albonotatus VIMMER, 1932; albosignatus VIMMER, 1932;

polymaculatus VIMMER, 1932; pulcher ZILAHI-SEBESS, 1934; kirovabadicus DZHAFAROV 1964;

matsuensis LIEN, WENG & LIN, 1996; meridionalis XUE, LIU & YU, 2003

- manchuriensis TOKUNAGA, 1941 (Culicoides)

Syn.: setiger GOETGHEBUER, 1938; goetghebueri ARNAUD, 1956; machardyi CAMPBELL & PELHAM-

CLINTON, 1960; ochraceimaculatus SHEVCHENKO, 1970; ochraceipennis SHEVCHENKO, 1970:

mesostigma REMM, 1971; vistulensis SKIERSKA, 1973

- salinarius KIEFFER, 1914 (Culicoides)

Syn.: halobius KIEFFER, 1914; meinerti KIEFFER, 1915; punctatidorsum KIEFFER, 1924

- sphagnumensis WILLIAMS, 1955 (Culicoides)

Syn.: lacicola ARNAUD, 1956; carjalaensis GLUKHOVA, 1957

Culicoides (Culicoides LATREILLE, 1809) (6)

- delta EDWARDS, in EDWARDS et AL. 1939 (Culicoides)

Syn.: lupicaris DOWNES & KETTLE, 1952

- flavipulicaris DZHAFAROV, 1964 (Culicoides)

- grisescens EDWARDS, in EDWARDS et AL. 1939 (Culicoides)

Syn.: remmi DAMIAN-GEORGESCU, 1972; arschanicus MIRZAEVA, 1984

- impunctatus GOETGHEBUER, 1920 (Culicoides)

Syn.: minor TOKUNAGA, 1941

- pulicaris (LINNAEUS, 1758) (Culex)

Syn.: setosinervis KIEFFER, 1913; pullatus KIEFFER, 1915; stephensi CARTER, 1916; cinerellus KIEFFER,

1919; quinquepunctatus GOETGHEBUER, 1921; flaviplumus KIEFFER, 1924; sawamotoi KONO &

TAKAHASI, 1940

- punctatus (MEIGEN, 1804) (Ceratopogon)

Syn.: punctatus LATREILLE, 1809; ocellaris KIEFFER, 1921; kasachstanicus SHAKIRZJANOVA, 1963

Culicoides (Monoculicoides KHALAF, 1954) (6)

- longicollis GLUKHOVA, 1971 (Culicoides)

- nubeculosus (MEIGEN, 1830) (Ceratopogon)

Syn.: puncticollis GOETGHEBUER, 1912; punctaticollis GOETGHEBUER, 1920

- parroti KIEFFER, 1922 (Culicoides)

- puncticollis (BECKER, 1903) (Ceratopogon)

Syn.: algecirensis STROBL, 1900; impressus KIEFFER, 1918; distigma KIEFFER, 1922; donatieni

KIEFFER, 1922; sciniphes KIEFFER, 1925; bipunctatus VIMMER, 1932; tripunctatus VIMMER, 1932;

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wenigi VIMMER, 1932; flavitarsis VIMMER, 1932; griseovittatus VIMMER, 1932; luteosignatus VIMMER,

1932; vavrai VIMMER, 1932

- riethi KIEFFER, 1914 (Culicoides)

Syn.: cordatus KIEFFER, 1921; crassiforceps KIEFFER, 1924; gigas ROOT & HOFFMAN, 1937

- stigma (MEIGEN, 1818) (Ceratopogon)

Syn.: kiefferi GOETGHEBUER, 1910; cordiformitarsis CARTER, 1916; unimaculatus GOETGHEBUER,

1920; stigmoides CALLOT, KREMER & DEDUIT, 1962

Culicoides (Oecacta POEY, 1851) (4)

- albicans (WINNERTZ, 1852) (Ceratopogon)

- clintoni BOORMAN, 1984 (Culicoides)

- semimaculatus CLASTRIER, 1958 (Culicoides)

Syn.: karajevi DZHAFAROV, 1961

- vexans (STAEGER, 1839) (Ceratopogon)

Syn.: pungens KIEFFER, 1901; perpungens KIEFFER, 1919; ajbassovi SHAKIRZJANOVA, 1962

Culicoides (Pontoculicoides REMM, 1968) (2)

- slovacus ORSZÁGH, 1969 (Culicoides)

- tauricus GUTSEVICH, 1959 (Culicoides)

Culicoides (Sensiculicoides SHEVCHENKO, 1977) (15)

- clastrieri CALLOT, KREMER & DEDUIT, 1962 (Culicoides)

- comosioculatus TOKUNAGA, 1956 (Culicoides)

Syn.: chaetophthalmus AMOSOVA, 1957; caucasicus SERGEJEV, 1959; setosus GUTSEVICH, 1960

- duddingstoni KETTLE & LAWSON, 1955 (Culicoides)

- festivipennis KIEFFER, 1914 (Culicoides)

Syn.: odibilis AUSTEN, 1921; winnertzi EDWARDS, 1926

- griseidorsum KIEFFER, 1918 (Culicoides)

- heliophilus EDWARDS, 1921 (Culicoides)

Syn.: latifrontis SHAKIRZJANOVA, 1962; kobachidzei DZHAFAROV, 1964

- jurensis CALLOT, KREMER & DEDUIT, 1962 (Culicoides)

- kibunensis TOKUNAGA, 1937 (Culicoides)

Syn.: cubitalis EDWARDS, in EDWARDS et AL. 1939; ponkikiri KONO & TAKAHASI, 1940; sitinohensis

OKADA, 1941

- maritimus KIEFFER, 1924 (Culicoides)

Syn.: submaritimus DZHAFAROV, 1962

- pictipennis (STAEGER, 1839) (Ceratopogon)

Syn.: arcuatus WINNERTZ, 1852; guttularis KIEFFER, 1919; maculatus ZILAHI-SEBESS, 1936;

achkamalicus DZHAFAROV, 1964; luganicus SHEVCHENKO, 1972

- pseudoheliophilus CALLOT & KREMER, 1961 (Culicoides)

Syn.: albihalteratus GOETGHEBUER, 1935

- simulator EDWARDS, in EDWARDS et AL. 1939 (Culicoides)

- ustinovi SCHEVCHENKO, 1962 (Culicoides)

- vidourlensis CALLOT, KREMER, MOLET & BACH, 1968 (Culicoides)

- zhogolevi REMM, in REMM & ZHOGOLEV 1968 (Culicoides)

Culicoides (Silvaticulicoides GLUKHOVA, 1977) (4)

- achrayi KETTLE & LAWSON, 1955 (Culicoides)

- fascipennis (STAEGER, 1839) (Ceratopogon)

86

Syn.: distictus KIEFFER, in THIENEMANN & KIEFFER 1916; albonotatus KIEFFER, 1918; turficola

KIEFFER, 1925

- pallidicornis KIEFFER, 1919 (Culicoides)

Syn.: susae KIEFFER, 1919; dileucus KIEFFER, 1921; brunneiscutellatus ZILAHI-SEBESS, 1933;

bruneoscutellatus ZILAHI-SEBESS, 1934; niger ROOT & HOFFMAN, 1937

- subfasciipennis KIEFFER, 1919 (Culicoides)

Syn.: analis KIEFFER, 1925

Culicoides (Wirthomyia VARGAS, 1973) (3)

- minutissimus (ZETTERSTEDT, 1855) (Ceratopogon)

Syn.: nanulus KIEFFER, 1919; albihalter KIEFFER, 1919; bychowskyi DZHAFAROV, 1964; tugaicus

DZHAFAROV, 1960

- reconditus CAMPBELL & PELHAM CLINTON, 1960 (Culicoides)

- segnis CAMPBELL & PELHAM CLINTON, 1960 (Culicoides)

Unplaced species (2)

- cameroni CAMPBELL & PELHAM CLINTON, 1960 (Culicoides)

- furcillatus CALLOT, KREMER & PARADIS, 1962 (Culicoides)

Tribe Ceratopogonini NEWMAN, 1834

Allohelea KIEFFER, 1917 (1)

- tessellata (ZETTERSTEDT, 1850) (Ceratopogon)

Syn.: illustris WINNERTZ, 1852

Alluaudomyia KIEFFER, 1913 (3)

- quadripunctata (GOETGHEBUER, 1934) (Culicoides)

- splendida (WINNERTZ, 1852) (Ceratopogon)

- wyskokensis SZADZIEWSKI & DOMINIAK, 2015 (Alluaudomyia)

Brachypogon KIEFFER, 1899 (14)

Brachypogon (Brachypogon KIEFFER, 1899) (3)

- beaufortanensis DELÉCOLLE & RIEB, 1992 (Brachypogon)

Syn.: sudowicus SZADZIEWSKI, 2001

- nieves (HAVELKA, 1976) (Ceratopogon)

- vitiosus (WINNERTZ, 1852) (Ceratopogon)

Syn.: minima KIEFFER, 1924; niger MAYER, 1934

Brachypogon (Isohelea KIEFFER, 1917) (11)

- babiogorensis SZADZIEWSKI, in SZADZIEWSKI et AL. 1994 (Brachypogon)

- beskidicus Krzywiński, in SZADZIEWSKI et AL. 1994 (Brachypogon)

- bialoviesicus KRZYWIŃSKI, in SZADZIEWSKI et AL. 1994 (Brachypogon)

- carpaticus SZADZIEWSKI, in SZADZIEWSKI et AL. 1994 (Brachypogon)

- hudjakovi (REMM, 1974) (Ceratopogon)

- incompletus (KIEFFER, 1925) (Trishelea)

Syn.: thienemanni MAYER, 1940; lapiae CLASTRIER, 1961; xuguitensis CAO & CHEN, 1984

- nitidulus (EDWARDS, 1921) (Psilohelea)

Syn.: crassiforceps KIEFFER, 1925; turfacea KIEFFER, 1925; finniae CLASTRIER, 1961

- norvegicus SZADZIEWSKI & HAGAN, 2000 (Brachypogon)

- silecis SZADZIEWSKI, 1990 (Brachypogon)

87

- sociabilis (GOETGHEBUER, 1920) (Psilohelea)

- zavoicus SZADZIEWSKI, in SZADZIEWSKI et AL. 1994 (Brachypogon)

Ceratoculicoides WIRTH & RATANAWORABHAN, 1971 (1)

- moravicus KNOZ, 1987 (Ceratoculicoides)

Syn.: gracilipes REMM, 1967; havelkai WIRTH & GROGAN, 1988; remmi GOSSERIES, 1989

Ceratopogon MEIGEN, 1803 (4)

- grandiforceps (KIEFFER, 1913) (Dasyhelea)

Syn.: parvula KIEFFER, 1925

- lacteipennis ZETTERSTEDT, 1838 (Ceratopogon)

Syn.: edentata EDWARDS, 1921; sphagnicola KIEFFER, 1924; conjuncta KIEFFER, 1925; longitarsis

MAYER, 1940

- naccinervis BORKENT, in BORKENT & WIRTH 1997 (Ceratopogon)

Syn.: crassinervis GOETGHEBUER, 1920 (preocc.)

- romanicus DAMIAN-GEORGESCU, 1972 (Ceratopogon)

Kolenohelea MEILLON & WIRTH, 1981 (1)

- calcarata (GOETGHEBUER, 1920) (Monohelea)

Syn.: sharpi EDWARDS, 1929; fusca GOETGHEBUER, 1932

Monohelea KIEFFER, 1917 (1)

- estonica REMM, 1965 (Monohelea)

Schizohelea KIEFFER, 1917 (1)

- leucopeza (MEIGEN, 1804) (Ceratopogon)

Syn.: albitarsis WIEDEMANN, 1817; copiosus WINNERTZ, 1852; politus COQUILLETT, 1901; stecki

KIEFFER, 1915; gorana MAYER, 1937; xanthopeza CLASTRIER, 1963

Serromyia MEIGEN, 1818 (8)

- atra (MEIGEN, 1818) (Ceratopogon)

Syn.: albitarsis KIEFFER, 1919; micronyx KIEFFER, 1919; spinosipes KIEFFER, 1919; nitens

GOETGHEBUER, 1920

- bicolor BORKENT, in BORKENT & BISSETT 1990 (Serromyia)

- femorata (MEIGEN, 1804) (Ceratopogon)

Syn.: armatus MEIGEN, 1818; foersteri MEIGEN, 1838; flavipes GIMMERTHAL, 1847; flavicrus KIEFFER,

1906; inermipes KIEFFER, 1919

- ledicola KIEFFER, 1925 (Serromyia)

Syn.: europaea CLASTRIER, 1963; macronyx GOETGHEBUER, 1933

- morio (FABRICIUS, 1775) (Culex)

Syn.: nudipennis KIEFFER, 1913

- rufitarsis (MEIGEN, 1818) (Ceratopogon)

Syn.: gelida KIEFFER, 1925; bispinosa GOETGHEBUER, 1936; dipetala REMM, 1965

- subinermis KIEFFER, 1919 (Serromyia)

- tecta BORKENT, in BORKENT & BISSETT 1990 (Serromyia)

88

Stilobezzia KIEFFER, 1911 (3)

Stilobezzia (Acanthohelea KIEFFER, 1917) (2)

- gracilis (HALIDAY, 1833) (Ceratopogon)

Syn.: dorsalis ZETTERSTEDT, 1850

- ochracea (WINNERTZ, 1852) (Ceratopogon)

Syn.: rufithorax KIEFFER, 1919; scutellata GOETGHEBUER, 1920

Stilobezzia (Stilobezzia KIEFFER, 1911) (1)

- antennalis (COQUILLETT, 1901) (Ceratopogon)

Tribe Heteromyiini WIRTH, 1962

Clinohelea KIEFFER, 1917 (1)

- unimaculata (MACQUART, 1826) (Ceratopogon)

Syn.: maculipes MEIGEN, 1830; variegatus WINNERTZ, 1852

Tribe Sphaeromiini NEWMAN, 1834

Macropeza MEIGEN, 1818 (1)

- albitarsis MEIGEN, 1818 (Macropeza)

Syn.: valvatus WINNERTZ, 1852

Mallochohelea WIRTH, 1962 (4)

- inermis (KIEFFER, 1909) (Johannseniella)

Syn.: kiefferi GOETGHEBUER, 1921

- munda (LOEW, 1864) (Ceratopogon)

Syn.: breviforceps KIEFFER, 1915; dentata KIEFFER, 1909

- nitida (MACQUART, 1826) (Ceratopogon)

Syn.: clavicornis KIEFFER, 1919; longiforceps KIEFFER, 1919

- setigera (LOEW, 1864) (Ceratopogon)

Syn.: maihensis REMM, 1971

Probezzia KIEFFER, 1906 (1)

- seminigra (PANZER, 1798) (Tipula)

Syn.: varipes CURTIS, 1829; munda LOEW, 1869; venustus MEIGEN, 1818; inflata WINNERTZ, 1852;

spinosidorsum KIEFFER, 1921; borealis CLASTRIER, 1962

Sphaeromias CURTIS, 1829 (2)

- fasciatus (MEIGEN, 1804) (Ceratopogon)

Syn.: cingulatus MEIGEN, 1804; albomarginatus CURTIS, 1829; procerus ZETTERSTEDT, 1855; ocularis

KIEFFER, 1924; goetghebueri DE MEIJERE, 1946

- pictus (MEIGEN, 1818) (Ceratopogon)

Syn.: punctatus MEIGEN, 1830; elegans WINNERTZ, 1852; candidatus LOEW, 1856; miricornis KIEFFER,

1919; copiosa KIEFFER, 1925; sparus BORKENT, in BORKENT & WIRTH 1997

Tribe Palpomyiini ENDERLEIN, 1836

Bezzia KIEFFER, 1899 (17)

- affinis (STAEGER, 1839) (Ceratopogon)

- albicornis (MEIGEN, 1818) (Ceratopogon)

89

Syn.: pallidetarsatus STROBL, 1900; strobli KIEFFER, 1919; brevinervis KIEFFER, 1919; atrata MACFIE,

1944

- annulipes (MEIGEN, 1830) (Ceratopogon)

Syn.: bidentata KIEFFER, 1901; fossicola KIEFFER, 1912; ploenensis KIEFFER, 1921; media KIEFFER,

1925; phragmitis KIEFFER, 1925; digramma KIEFFER, 1925; kyotoensis TOKUNAGA, 1939; sicarti

CLASTRIER, 1962

- bicolor (MEIGEN, 1804) (Ceratopogon)

Syn.: typhae KIEFFER, 1919; brehmiana KIEFFER, 1924; algarum ZILAHI-SEBESS, 1930; rufithorax

GOETGHEBUER, 1933; copiosa THOMSEN, 1935

- circumdata (STAEGER, 1839) (Ceratopogon)

Syn.: solstitialis WINNERTZ, 1852; hydrophila KIEFFER, 1909; sieberti KIEFFER, 1921; aquatilis

GOETGHEBUER, 1923; stagnalis KIEFFER, 1925

- coracina (ZETTERSTEDT, 1850) (Ceratopogon)

Syn.: albipes WINNERTZ, 1852; fusciclava KIEFFER, 1919

- elongata ZILAHI-SEBESS, 1940 (Bezzia)

- flavicornis (STAEGER, 1839) (Ceratopogon)

Syn.: flavipalpis WINNERTZ, 1852; flavipluma KIEFFER, 1919; spinifera GOETGHEBUER, 1920;

spinosula CLASTRIER, 1962

- kazlauskasi REMM, 1966 (Bezzia)

- leucogaster (ZETTERSTEDT, 1850) (Ceratopogon)

Syn.: xanthocephala GOETGHEBUER, 1911; parvidens KIEFFER, 1914; picticornis KIEFFER, 1914;

danica KIEFFER, 1915; grisea KIEFFER, 1919; brachycera KIEFFER, 1919; belgica KIEFFER, 1919;

hungarica ZILAHI-SEBESS, 1930

- nigrita CLASTRIER, 1962 (Bezzia)

- nigritula (ZETTERSTEDT, 1838) (Ceratopogon)

Syn.: tenebricosa GOETGHEBUER, 1912; atripes ZILAHI-SEBESS, 1940

- nobilis (WINNERTZ, 1852) (Ceratopogon)

Syn.: setulosus LOEW, 1861; barberi COQUILLETT, 1901; punctata LUNDSTRÖM, 1910; leucosticta

KIEFFER, 1919; cinerella KIEFFER, 1919; armaticrus KIEFFER, 1919; csikiana KIEFFER, 1919;

chrysocoma KIEFFER, 1922; acanthodes MACFIE, 1940; atlantica WIRTH & WILLIAMS, 1957;

nobiliformis CLASTRIER, 1962

- ornata (MEIGEN, 1830) (Ceratopogon)

Syn.: vittiger ZETTERSTEDT, 1850; albosignata KIEFFER, 1919

- rhynchostylata REMM, 1974 (Bezzia)

- signata (MEIGEN, 1804) (Ceratopogon)

Syn.: heusdensis GOETGHEBUER, 1935

- winnertziana KIEFFER, 1919 (Bezzia)

Syn.: gracilis WINNERTZ, 1852

Palpomyia MEIGEN, 1818 (13)

- armipes (MEIGEN, 1838) (Ceratopogon)

Syn.: erythrocephala STAEGER, 1839; rufipectus WINNERTZ, 1852

- brachialis (HALIDAY, 1833) (Ceratopogon)

Syn.: longipennis KIEFFER, 1919; fusciclava KIEFFER, 1919; nemorivaga GOETGHEBUER, 1920

- distincta (HALIDAY, 1833) (Ceratopogon)

90

Syn.: ephippium ZETTERSTEDT, 1855; rubra KIEFFER, 1919; flaviscutellum KIEFFER, 1919; ruficornis

GOETGHEBUER, 1934

- flavipes (MEIGEN, 1804) (Ceratopogon)

Syn.: hortulanus MEIGEN, 1818; mutabilis CLASTRIER, 1962

- infuscata KIEFFER, 1919 (Palpomyia)

- lineata (MEIGEN, 1804) (Ceratopogon)

Syn.: cinerea MACQUART, 1826; binotata STAEGER, 1839; slossonae COQUILLETT, 1905; algarum

KIEFFER, in KIEFFER & THIENEMANN 1908; winnertzi KIEFFER, 1914; illinoisensis MALLOCH, 1914;

octasema KIEFFER, 1914; spinosissima KIEFFER, 1915; unifasciata KIEFFER, 1922; murina KIEFFER,

1925; obesa GOETGHEBUER, 1927; nigrotibialis GOETGHEBUER, 1933; pruinescens THOMSEN, 1935;

edwardsi DE MEIJERE, 1946; stagnalis CLASTRIER, 1962

- nigripes (MEIGEN, 1830) (Ceratopogon)

Syn.: pratensis MEIGEN, 1830; bryocrypta KIEFFER, 1925

- praeusta (LOEW, 1869) (Ceratopogon)

Syn.: apicalis STROBL, 1898; terminalis KIEFFER, 1919; schineri GOETGHEBUER, 1920

- rufipes (MEIGEN, 1818) (Ceratopogon)

Syn.: spinosior KIEFFER, 1919; pseudospinipes GOETGHEBUER, 1933; plurispinosa CLASTRIER, 1962

- serripes (MEIGEN, 1818) (Ceratopogon)

Syn.: transfuga STAEGER, 1839; tarsata ZETTERSTEDT, 1855; ruficeps KIEFFER, 1918

- spinipes (MEIGEN, in PANZER 1806) (Ceratopogon)

Syn.: fulva MACQUART, 1826; ferrugineus MEIGEN, 1830; parviforceps KIEFFER, 1925

- tibialis (MEIGEN, 1818) (Ceratopogon)

Syn.: atripecta KIEFFER, 1919; laticollis GOETGHEBUER, 1922; nipponica TOKUNAGA, 1939

- tinctipennis KIEFFER, 1919 (Palpomyia)

Syn.: obscuripennis KIEFFER, 1919; semifumosa GOETGHEBUER, 1922; adusta KIEFFER, 1925; nigritella

CLASTRIER, 1962

Phaenobezzia Haeselbarth, 1965 (1)

- rubiginosa (WINNERTZ, 1852) (Ceratopogon)

Syn.: glyceriae KIEFFER, 1913; nitidiventris GOETGHEBUER, 1923; radialis GOETGHEBUER, 1936

NEW RECORD

Ceratopogon naccinervis BORKENT, in BORKENT & WIRTH 1997

Material examined. Poland, Kampinoski Park Narodowy nr Warsaw, grąd-wiąz (Ulmus), 16-

21 May 1979, yellow pan trap, IZPAN, 2 males, deposited in the coll. of University of

Gdańsk, CEIUG.

Comments. Males of the species are characteristic in having short gonocoxite, long and

slender gonostylus armed with apical 2 or 3 teeth (Fig. A), apicolateral process of tergite IX

long, slender and armed with 1-2 apical setae, parameres distinctly expanded in the distal half,

apex pointed, aedeagus with lateral projections bearing some apical teeth (Fig. A-D). Wing

length 1.30-1.32 mm, gonocoxite length 0.25-0.26 mm, gonocoxite length/head width 0.63,

gonocoxite about 2.5 times longer than broad. Ceratopogon naccinervis belongs to the group

of four very close species inluding also Nearctic C. abstrusus BORKENT & GROGAN, 1995, C.

91

arcanus BORKENT & GROGAN, 1995, and C. inverecundus BORKENT & GROGAN, 1995. Male

genitalia of Polish specimens are exactly the same as those of C. crassinervis from Estonia

(REMM 1974) and C. inverecundus from North America (BORKENT & GROGAN 1995). The

species is reported from Belgium, Estonia and Turkey (BORKENT & GROGAN 1995). From

Poland recorded for the first time.

FIG. Ceratopogon naccinervis BORKENT, in BORKENT & WIRTH 1997: male genitalia. A – ventral

aspect, aedeagus removed, B – apicolateral processes of tergite IX, C – aedeagus, D – parameres

ACKNOWLEGEMENTS

We are garteful to Dr. ELŻBIETA SONTAG (Department of Invertebrate Zoology

and Parasitology, University of Gdańsk, Poland) for preparing the plate with figures

of Ceratopogon naccinervis. Our thanks are due to Dr. JOANNA CHOUFANI (Department

of Sciences, Notre Dame University, Lebanon) and an anonymous Reviewer for valuable

comments that helped us to improve this paper.

92

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